Birds evolved from certain
featheredtheropod dinosaurs, and there is no real dividing line between birds and
non-avian dinosaurs except that some of the former survived the
Cretaceous–Paleogene extinction event while the latter did not. For the purposes of this article, a 'bird' is considered to be any member of the clade
Aves in the broadest sense.[1] Some dinosaur groups which may or may not be true birds are listed below under
Proto-birds.
This page contains a listing of prehistoric bird taxa only known from completely
fossilized specimens. These extinctions took place before the Late
Quaternary and thus took place in the absence of significant human interference. While the earliest
hominids had been eating birds and especially their eggs,
human population and
technology was simply insufficient to seriously affect healthy bird populations until the
Upper Paleolithic Revolution. Rather, reasons for the extinctions listed here are stochastic abiotic events such as
bolide impacts,
climate changes due to orbital shifts, mass volcanic eruptions etc. Alternatively, species may have gone extinct due to
evolutionary displacement by successor or competitor taxa – it is notable that an extremely large number of seabirds have gone extinct during the mid-
Tertiary; this seems at least partly due to competition by the contemporary radiation of
marine mammals.
The relationships of these
taxa are often hard to determine, as many are known only from very fragmentary remains and due to the complete fossilization precluding analysis of information from
DNA,
RNA or protein sequencing. The taxa listed in this article should be classified with the
Wikipedia conservation status category "Fossil".
Before the late 19th century, when
minerals were still considered one of the
kingdoms of
binomial nomenclature, fossils were often treated according to a parallel
taxonomy. Rather than assigning them to animal or plant genera, they were treated as mineral genera and given
binomial names typically using Osteornis ("bone-bird") or Ornitholithus ("bird fossil") as "
genus". The latter name, however, is still in use for an
oogenus of
fossil bird eggs. Also, other animals (in particular
pterosaurs) were placed in these "genera". In sources pre-dating the
Linnean system, the above terms are also seen in the more extensive descriptions used to name
taxa back then.
Taxonomic list of fossil prehistoric birds
Higher-level
taxa are presented in likely or suspected
phylogenetic order.
Genus-level taxa and lower are sorted chronologically, in ascending order (i.e., older taxa first).
The higher-level groups of non-
Neornithes are arranged based on the
phylogeny proposed by
Luis Chiappe,[2] updated and expanded to incorporate recent research.[3] The categories are inclusive in ascending order.
Taxonomic assignments, especially in the pygostylian to early neornithine genera, are still very provisional and subject to quite frequent change.[4][5]
The most primitive "birds", usually still possessing a long bony tail with generally unfused vertebrae. Not all of these may be on the line of bird ancestors; whether they are not closer to other
theropods groups than to the
Avialae remains to be thoroughly tested (see Xiaotingia).
The earliest birds with a modern
pygostyle: a reduction and fusion of the tail vertebrae; possibly a
paraphyletic group. Two types of pygostyle are known, a rod-shaped one found in
Confuciusornithidae,
Enantiornithes and some non-avian theropods such as Nomingia, and a
plowshare-shaped one, only known in the lineage leading to modern birds. It is not certain that the pygostyles found in birds are indeed
synapomorphies.[6]
Note that Holtz (2011) also included Zhyraornis in his classification of euenantiornithines, though this genus is more often classified as an
ornithuran.[13] Holtz also placed Liaoningornis as an
ornithuromorph, though more recent studies have placed it as a close relative of Eoalulavis.[14]
Also called "basal
Ornithuromorpha".[15] Essentially modern birds, except many still possess a few primitive features such as teeth or wing claws. These have the
plowshare-shaped pygostyle and proper tail fan as seen in most living birds. The
taxonomy of this group is confusing; the name "Ornithurae" was first proposed by
Ernst Haeckel in 1866 and has been revised in meaning several times since.
The following is a list of primitive euornithian genera and those that cannot be confidently referred to any subgroups, following Holtz (2011).[7]
Unresolved and basal forms
These modern birds are known from remains that cannot be placed in relation to any one modern group and are neither
autapomorphic enough to assign them to own orders. Especially the
Late Cretaceous/early
Paleogenetaxa are probably
basal to several modern orders, while later
Paleogene taxa often represent extinct lineages outside the modern families.
†"Palaeotringa" vetusMarsh 1870 (Lance Late Cretaceous of Wyoming – Hornerstown Late Cretaceous/?Early Palaeocene of New Jersey, US) - gruiform? anseriform (presbyornithid)?
†Teleornis impressusAmeghino 1899 (Deseado Early Oligocene of Argentina)– anatid?
†Pseudolarus guaraniticusAmeghino 1899 (Deseado Early Oligocene – Miocene of Argentina) – gruiform?
Neornithes incerta sedis BMNH PAL 4989 (Hampstead Early Oligocene of Isle of Wight, England) – formerly "
Ptenornis" and included in Headonornis; anseriform?
†"Anas" creccoidesvan Beneden 1871 (Early-mid Oligocene of Belgium) – anseriform?
The diatrymas, a group of huge flightless Paleogene birds of unclear affinities. Traditionally placed within the Gruiformes, they are usually considered a distinct order nowadays and appear closer to the Anseriformes.
The group that includes modern
pelicans and
cormorants. As presented here
paraphyletic; the tropicbird lineage is not part of this group and relationships with Procellariiformes and Sphenisciformes require more research. Also, as the pelicans are at least as close to the
Ciconiiformes as to cormorants, the latter group is being recognized as Phalacrocoraciiformes by some recent authors and the core Pelecaniformes are occasionally merged into the Ciconiiformes.
†Rupelornis van Beneden 1871 [GaviotaFischer 1983 non Miller & Sibley 1941; ArdeitaCheneval 1996; DiomedeoidesFischer 1985; FrigidafronsCheneval 1995] (Early Oligocene– Early Miocene of C Europe and Iran)
These fossil taxa cannot be assigned to any major group with reasonable certainty. The "proto-birds" above are of some indeterminate basal position in the entire avialan (and paravian) radiation, but known from such diagnostic material that their relationships at the
family level are known. In contrast, the taxa here have a
hypodigm that is usually just sufficient for giving them a valid scientific name, but not for phylogenetic purposes beyond classing them as pygostylians or more modern birds. Some, however, are known from such fragmentary remains that the possibility that they are non-avian "
reptiles" such as
dinosaurs cannot be ruled out at present.
†"Ichthyornis" minusculus Nesov 1990 (Bissekty Late Cretaceous of Kyzyl Kum, Uzbekistan) – enantiornithine?
^Brodkob, Pierce (1963). "Catalogue of fossil birds 1- Archaeopterygiformes through Ardeiformes". Biological Sciences, Bulletin of the Florida State Museum. 7 (4): 180–293.
^O'Connor, J. K.; Zhang, Y.; Chiappe, L. M.; Meng, Q.; Quanguo, L.; Di, L. (2013). "A new enantiornithine from the Yixian Formation with the first recognized avian enamel specialization". Journal of Vertebrate Paleontology. 33 (1): 1–12.
Bibcode:
2013JVPal..33....1O.
doi:
10.1080/02724634.2012.719176.
S2CID85261944.
^Gareth J. Dyke, Attila Ősi (2010). "A review of Late Cretaceous fossil birds from Hungary". Geological Journal. 45 (4): 434–444.
doi:
10.1002/gj.1209.
S2CID128580503.
^
abNikita V. Zelenkov; Thomas A. Stidham; Nicolay V. Martynovich; Natalia V. Volkova; Qiang Li; Zhuding Qiu (2018). "The middle Miocene duck Chenoanas (Aves, Anatidae): new species, phylogeny and geographical range". Papers in Palaeontology. 4 (3): 309–326.
doi:
10.1002/spp2.1107.
S2CID134072594.
^Similar to Urocolius and Limnatornis (if distinct): Mlíkovský (2002)
References
Chiappe, Luis M. (2001): The rise of birds. In:
Briggs, Derek E.G. & Crowther, P.R. (eds.): Palaeobiology II: A Synthesis: 102-106. Cambridge University Press, Cambridge, UK.
Chiappe, Luis M. (2002): Basal bird phylogeny: problems and solutions. In: Chiappe, L.M. and Witmer, L.M. (eds.): Mesozoic Birds: Above the Heads of Dinosaurs: 448-472. University of California Press, Berkeley, US.
Gauthier, Jacques A. & de Queiroz, Kevin (2001): Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the name "Aves". In: Gauthier, Jacques & Gall, L.F. (eds.): New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom: 7-41. Peabody Museum of Natural History, Yale University, New Haven, Connecticut.
Mortimer, Michael (2004): The Theropod Database:
Phylogeny of taxa. Retrieved 2013-MAR-02.
Olson, Storrs L. (1985): The fossil record of birds. In: Farner, D.S.; King, J.R. & Parkes, Kenneth C. (eds.): Avian Biology8: 79-238. Academic Press, New York. Not in copyright;
PDF fulltext
Birds evolved from certain
featheredtheropod dinosaurs, and there is no real dividing line between birds and
non-avian dinosaurs except that some of the former survived the
Cretaceous–Paleogene extinction event while the latter did not. For the purposes of this article, a 'bird' is considered to be any member of the clade
Aves in the broadest sense.[1] Some dinosaur groups which may or may not be true birds are listed below under
Proto-birds.
This page contains a listing of prehistoric bird taxa only known from completely
fossilized specimens. These extinctions took place before the Late
Quaternary and thus took place in the absence of significant human interference. While the earliest
hominids had been eating birds and especially their eggs,
human population and
technology was simply insufficient to seriously affect healthy bird populations until the
Upper Paleolithic Revolution. Rather, reasons for the extinctions listed here are stochastic abiotic events such as
bolide impacts,
climate changes due to orbital shifts, mass volcanic eruptions etc. Alternatively, species may have gone extinct due to
evolutionary displacement by successor or competitor taxa – it is notable that an extremely large number of seabirds have gone extinct during the mid-
Tertiary; this seems at least partly due to competition by the contemporary radiation of
marine mammals.
The relationships of these
taxa are often hard to determine, as many are known only from very fragmentary remains and due to the complete fossilization precluding analysis of information from
DNA,
RNA or protein sequencing. The taxa listed in this article should be classified with the
Wikipedia conservation status category "Fossil".
Before the late 19th century, when
minerals were still considered one of the
kingdoms of
binomial nomenclature, fossils were often treated according to a parallel
taxonomy. Rather than assigning them to animal or plant genera, they were treated as mineral genera and given
binomial names typically using Osteornis ("bone-bird") or Ornitholithus ("bird fossil") as "
genus". The latter name, however, is still in use for an
oogenus of
fossil bird eggs. Also, other animals (in particular
pterosaurs) were placed in these "genera". In sources pre-dating the
Linnean system, the above terms are also seen in the more extensive descriptions used to name
taxa back then.
Taxonomic list of fossil prehistoric birds
Higher-level
taxa are presented in likely or suspected
phylogenetic order.
Genus-level taxa and lower are sorted chronologically, in ascending order (i.e., older taxa first).
The higher-level groups of non-
Neornithes are arranged based on the
phylogeny proposed by
Luis Chiappe,[2] updated and expanded to incorporate recent research.[3] The categories are inclusive in ascending order.
Taxonomic assignments, especially in the pygostylian to early neornithine genera, are still very provisional and subject to quite frequent change.[4][5]
The most primitive "birds", usually still possessing a long bony tail with generally unfused vertebrae. Not all of these may be on the line of bird ancestors; whether they are not closer to other
theropods groups than to the
Avialae remains to be thoroughly tested (see Xiaotingia).
The earliest birds with a modern
pygostyle: a reduction and fusion of the tail vertebrae; possibly a
paraphyletic group. Two types of pygostyle are known, a rod-shaped one found in
Confuciusornithidae,
Enantiornithes and some non-avian theropods such as Nomingia, and a
plowshare-shaped one, only known in the lineage leading to modern birds. It is not certain that the pygostyles found in birds are indeed
synapomorphies.[6]
Note that Holtz (2011) also included Zhyraornis in his classification of euenantiornithines, though this genus is more often classified as an
ornithuran.[13] Holtz also placed Liaoningornis as an
ornithuromorph, though more recent studies have placed it as a close relative of Eoalulavis.[14]
Also called "basal
Ornithuromorpha".[15] Essentially modern birds, except many still possess a few primitive features such as teeth or wing claws. These have the
plowshare-shaped pygostyle and proper tail fan as seen in most living birds. The
taxonomy of this group is confusing; the name "Ornithurae" was first proposed by
Ernst Haeckel in 1866 and has been revised in meaning several times since.
The following is a list of primitive euornithian genera and those that cannot be confidently referred to any subgroups, following Holtz (2011).[7]
Unresolved and basal forms
These modern birds are known from remains that cannot be placed in relation to any one modern group and are neither
autapomorphic enough to assign them to own orders. Especially the
Late Cretaceous/early
Paleogenetaxa are probably
basal to several modern orders, while later
Paleogene taxa often represent extinct lineages outside the modern families.
†"Palaeotringa" vetusMarsh 1870 (Lance Late Cretaceous of Wyoming – Hornerstown Late Cretaceous/?Early Palaeocene of New Jersey, US) - gruiform? anseriform (presbyornithid)?
†Teleornis impressusAmeghino 1899 (Deseado Early Oligocene of Argentina)– anatid?
†Pseudolarus guaraniticusAmeghino 1899 (Deseado Early Oligocene – Miocene of Argentina) – gruiform?
Neornithes incerta sedis BMNH PAL 4989 (Hampstead Early Oligocene of Isle of Wight, England) – formerly "
Ptenornis" and included in Headonornis; anseriform?
†"Anas" creccoidesvan Beneden 1871 (Early-mid Oligocene of Belgium) – anseriform?
The diatrymas, a group of huge flightless Paleogene birds of unclear affinities. Traditionally placed within the Gruiformes, they are usually considered a distinct order nowadays and appear closer to the Anseriformes.
The group that includes modern
pelicans and
cormorants. As presented here
paraphyletic; the tropicbird lineage is not part of this group and relationships with Procellariiformes and Sphenisciformes require more research. Also, as the pelicans are at least as close to the
Ciconiiformes as to cormorants, the latter group is being recognized as Phalacrocoraciiformes by some recent authors and the core Pelecaniformes are occasionally merged into the Ciconiiformes.
†Rupelornis van Beneden 1871 [GaviotaFischer 1983 non Miller & Sibley 1941; ArdeitaCheneval 1996; DiomedeoidesFischer 1985; FrigidafronsCheneval 1995] (Early Oligocene– Early Miocene of C Europe and Iran)
These fossil taxa cannot be assigned to any major group with reasonable certainty. The "proto-birds" above are of some indeterminate basal position in the entire avialan (and paravian) radiation, but known from such diagnostic material that their relationships at the
family level are known. In contrast, the taxa here have a
hypodigm that is usually just sufficient for giving them a valid scientific name, but not for phylogenetic purposes beyond classing them as pygostylians or more modern birds. Some, however, are known from such fragmentary remains that the possibility that they are non-avian "
reptiles" such as
dinosaurs cannot be ruled out at present.
†"Ichthyornis" minusculus Nesov 1990 (Bissekty Late Cretaceous of Kyzyl Kum, Uzbekistan) – enantiornithine?
^Brodkob, Pierce (1963). "Catalogue of fossil birds 1- Archaeopterygiformes through Ardeiformes". Biological Sciences, Bulletin of the Florida State Museum. 7 (4): 180–293.
^O'Connor, J. K.; Zhang, Y.; Chiappe, L. M.; Meng, Q.; Quanguo, L.; Di, L. (2013). "A new enantiornithine from the Yixian Formation with the first recognized avian enamel specialization". Journal of Vertebrate Paleontology. 33 (1): 1–12.
Bibcode:
2013JVPal..33....1O.
doi:
10.1080/02724634.2012.719176.
S2CID85261944.
^Gareth J. Dyke, Attila Ősi (2010). "A review of Late Cretaceous fossil birds from Hungary". Geological Journal. 45 (4): 434–444.
doi:
10.1002/gj.1209.
S2CID128580503.
^
abNikita V. Zelenkov; Thomas A. Stidham; Nicolay V. Martynovich; Natalia V. Volkova; Qiang Li; Zhuding Qiu (2018). "The middle Miocene duck Chenoanas (Aves, Anatidae): new species, phylogeny and geographical range". Papers in Palaeontology. 4 (3): 309–326.
doi:
10.1002/spp2.1107.
S2CID134072594.
^Similar to Urocolius and Limnatornis (if distinct): Mlíkovský (2002)
References
Chiappe, Luis M. (2001): The rise of birds. In:
Briggs, Derek E.G. & Crowther, P.R. (eds.): Palaeobiology II: A Synthesis: 102-106. Cambridge University Press, Cambridge, UK.
Chiappe, Luis M. (2002): Basal bird phylogeny: problems and solutions. In: Chiappe, L.M. and Witmer, L.M. (eds.): Mesozoic Birds: Above the Heads of Dinosaurs: 448-472. University of California Press, Berkeley, US.
Gauthier, Jacques A. & de Queiroz, Kevin (2001): Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the name "Aves". In: Gauthier, Jacques & Gall, L.F. (eds.): New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom: 7-41. Peabody Museum of Natural History, Yale University, New Haven, Connecticut.
Mortimer, Michael (2004): The Theropod Database:
Phylogeny of taxa. Retrieved 2013-MAR-02.
Olson, Storrs L. (1985): The fossil record of birds. In: Farner, D.S.; King, J.R. & Parkes, Kenneth C. (eds.): Avian Biology8: 79-238. Academic Press, New York. Not in copyright;
PDF fulltext